Great Tit
Encyclopedia
The Great Tit is a passerine
Passerine
A passerine is a bird of the order Passeriformes, which includes more than half of all bird species. Sometimes known as perching birds or, less accurately, as songbirds, the passerines form one of the most diverse terrestrial vertebrate orders: with over 5,000 identified species, it has roughly...

 bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

 in the tit family Paridae. It is a widespread and common species throughout Europe, the Middle East, Central and Northern Asia, and parts of North Africa in any sort of woodland. It is generally resident, and most Great Tits do not migrate
Bird migration
Bird migration is the regular seasonal journey undertaken by many species of birds. Bird movements include those made in response to changes in food availability, habitat or weather. Sometimes, journeys are not termed "true migration" because they are irregular or in only one direction...

 except in extremely harsh winters. Until 2005 this species was lumped
Lumpers and splitters
Lumping and splitting refers to a well-known problem in any discipline which has to place individual examples into rigorously defined categories. The lumper/splitter problem occurs when there is the need to create classifications and assign examples to them, for example schools of literature,...

 with numerous other subspecies. DNA
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...

 studies have shown these other subspecies to be distinctive from the Great Tit and these have now been separated as two separate species, the Cinereous Tit
Cinereous Tit
The Cinereous Tit is a species of bird in the tit family Paridae. This species is made up of several populations that were earlier treated as subspecies of the Great Tit . These birds are grey backed with white undersides. The Great Tit in the new sense is distinguishable by the greenish-back and...

 of southern Asia, and the Japanese Tit
Japanese Tit
Japanese Tit , also known as Oriental Tit, is a passerine bird which replaces the similar Great Tit in Japan and the Russian Far East beyond the Amur River, including the Kuril Islands...

 of East Asia. The Great Tit remains the most widespread species in the genus Parus.

The Great Tit is a distinctive bird, with a black head and neck, prominent white cheeks, olive upperparts and yellow underparts, with some variation amongst the numerous subspecies. It is predominantly insectivorous
Insectivore
An insectivore is a type of carnivore with a diet that consists chiefly of insects and similar small creatures. An alternate term is entomophage, which also refers to the human practice of eating insects....

 in the summer, but will consume a wider range of food items in the winter months, including small hibernating bats. Like all tits it is a cavity nester, usually nesting in a hole in a tree. The female lays around 12 eggs and incubates them alone, although both parents raise the chicks. In most years the pair will raise two broods. The nests may be raided by woodpecker
Great Spotted Woodpecker
The Great Spotted Woodpecker , Dendrocopos major, is a bird species of the woodpecker family . It is distributed throughout Europe and northern Asia, and usually resident year-round except in the colder parts of its range...

s, squirrel
Eastern Gray Squirrel
The eastern gray squirrel is a tree squirrel in the genus Sciurus native to the eastern and midwestern United States, and to the southerly portions of the eastern provinces of Canada...

s and weasel
Least Weasel
The least weasel is the smallest member of the Mustelidae , native to Eurasia, North America and North Africa, though it has been introduced elsewhere. It is classed as Least Concern by the IUCN, due to its wide distribution and presumably large population...

s and infested with flea
Flea
Flea is the common name for insects of the order Siphonaptera which are wingless insects with mouthparts adapted for piercing skin and sucking blood...

s, and adults may be hunted by Sparrowhawks. The Great Tit has adapted well to human changes in the environment and is a common and familiar bird in urban parks and gardens. The Great Tit is also an important study species in ornithology
Ornithology
Ornithology is a branch of zoology that concerns the study of birds. Several aspects of ornithology differ from related disciplines, due partly to the high visibility and the aesthetic appeal of birds...

.

Taxonomy

The Great Tit was originally described under its current binomial name by Linnaeus in his 18th century work, Systema Naturae
Systema Naturae
The book was one of the major works of the Swedish botanist, zoologist and physician Carolus Linnaeus. The first edition was published in 1735...

. Its scientific name is derived from the Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

 parus "tit" and maior "larger".
The Great Tit was formerly treated as ranging from Britain to Japan and south to the islands of Indonesia, with 36 described subspecies ascribed to four main species groups. The major group had 13 subspecies across Europe, temperate Asia and north Africa, the minor group's nine subspecies occurred from southeast Russia and Japan into northern southeast Asia and the 11 subspecies in the cinereus group were found from Iran across south Asia to Indonesia. The three bokharensis subspecies were often treated as a separate species, Parus bokharensis, the Turkestan Tit. This form was once thought to form a ring species
Ring species
In biology, a ring species is a connected series of neighboring populations, each of which can interbreed with closely sited related populations, but for which there exist at least two "end" populations in the series, which are too distantly related to interbreed, though there is a potential gene...

 around the Tibetan Plateau
Tibetan Plateau
The Tibetan Plateau , also known as the Qinghai–Tibetan Plateau is a vast, elevated plateau in Central Asia covering most of the Tibet Autonomous Region and Qinghai, in addition to smaller portions of western Sichuan, southwestern Gansu, and northern Yunnan in Western China and Ladakh in...

, with gene flow throughout the subspecies, but this theory was abandoned when sequences of mitochondrial DNA
Mitochondrial DNA
Mitochondrial DNA is the DNA located in organelles called mitochondria, structures within eukaryotic cells that convert the chemical energy from food into a form that cells can use, adenosine triphosphate...

 were examined, finding that the four groups were distinct (monophyletic
Monophyly
In common cladistic usage, a monophyletic group is a taxon which forms a clade, meaning that it contains all the descendants of the possibly hypothetical closest common ancestor of the members of the group. The term is synonymous with the uncommon term holophyly...

) and that the hybridisation zones between the groups were the result of secondary contact after a temporary period of isolation.

A study published in 2005 confirmed that the major group was distinct from the cinereus and minor groups and that along with P.m. bokharensis it diverged from these two groups around 1.5 million years ago. The divergence between the bokharensis and major groups was estimated to have been about half a million years ago. The study also examined hybrids between representatives of the major and minor groups in the Amur Valley where the two meet. Hybrids were rare, suggesting that there were some reproductive barriers between the two groups. The study recommended that the two eastern groups be split out as new species, the Cinereous Tit
Cinereous Tit
The Cinereous Tit is a species of bird in the tit family Paridae. This species is made up of several populations that were earlier treated as subspecies of the Great Tit . These birds are grey backed with white undersides. The Great Tit in the new sense is distinguishable by the greenish-back and...

 (Parus cinereus), and the Japanese Tit
Japanese Tit
Japanese Tit , also known as Oriental Tit, is a passerine bird which replaces the similar Great Tit in Japan and the Russian Far East beyond the Amur River, including the Kuril Islands...

 (Parus minor), but that the Turkestan Tit be lumped
Lumpers and splitters
Lumping and splitting refers to a well-known problem in any discipline which has to place individual examples into rigorously defined categories. The lumper/splitter problem occurs when there is the need to create classifications and assign examples to them, for example schools of literature,...

 in with the Great Tit. This taxonomy has been followed by some authorities, for example the IOC World Bird List
Birds of the World: Recommended English Names
Birds of the World: Recommended English Names is a paperback book, written by Frank Gill and Minturn Wright on behalf of the International Ornithological Congress. The book is an attempt to produce a standardised set of English names for all bird species, and it is the product of a project set in...

. The Handbook of the Birds of the World
Handbook of the Birds of the World
The Handbook of the Birds of the World is a multi-volume series produced by the Spanish publishing house Lynx Edicions. It is the first handbook to cover every known living species of bird. The series is edited by Josep del Hoyo, Andrew Elliott, Jordi Sargatal and David A Christie.So far, 15...

volume treating the Parus species went for the more traditional classification, treating the Turkestan Tit as a separate species but retaining the Japanese and Cinereous Tits with the Great Tit, a move that has not been without criticism.

The nominate subspecies of the Great Tit is the most widespread, its range stretching from the Iberian Peninsula to the Amur Valley and from Scandinavia to the Middle East. The other subspecies have much more restricted distributions, four being restricted to islands and the remainder of the P. m. major subspecies representing former glacial refuge populations. The dominance of a single, morphologically uniform subspecies over such a large area suggests that the nominate race rapidly recolonised a large area after the last glacial epoch
Ice age
An ice age or, more precisely, glacial age, is a generic geological period of long-term reduction in the temperature of the Earth's surface and atmosphere, resulting in the presence or expansion of continental ice sheets, polar ice sheets and alpine glaciers...

. This hypothesis is supported by genetic studies which suggest a geologically recent genetic bottleneck
Population bottleneck
A population bottleneck is an evolutionary event in which a significant percentage of a population or species is killed or otherwise prevented from reproducing....

 followed by a rapid population expansion.

The genus Parus once held most of the species of tit in the family Paridae, but morphological and genetic studies led to the splitting of that large genus in 1998. The Great Tit was retained in Parus, which, along with Cyanistes
Cyanistes
Cyanistes is a genus of tits. It is frequently subsumed into Parus. It consists of three small species found in temperate Europe and Asia and Northern AfricaThe species are:* Blue Tit, Cyanistes caeruleus* Azure Tit, Cyanistes cyanus...

comprise a lineage of tits known as the "non-hoarders", with reference to the hoarding behaviour of members of the other clade. The genus Parus is still the largest in the family, but may be split again. Other than those species formerly considered to be subspecies, the Great Tit's closest relatives are the White-naped
White-winged Tit
The White-winged Tit sometimes called the White-naped Tit is a passerine bird in the tit family Paridae. It is endemic to India where it is found in dry thorn scrub forest in two disjunct populations, in western India and southern India...

 and Green-backed Tit
Green-backed Tit
The Green-backed Tit is a species of bird in the Paridae family.It is found in Bangladesh, Bhutan, China, India, Laos, Burma, Nepal, Pakistan, Taiwan, and Vietnam....

s of southern Asia. Hybrids with tits outside the Parus genus are very rare, but have been recorded with Blue Tit
Blue Tit
The Blue Tit is a 10.5 to 12 cm long passerine bird in the tit family Paridae. It is a widespread and common resident breeder throughout temperate and subarctic Europe and western Asia in deciduous or mixed woodlands...

, Coal Tit
Coal Tit
The Coal Tit, Periparus ater, is a passerine bird in the tit family Paridae. It is a widespread and common resident breeder throughout temperate to subtropical Eurasia and northern Africa. The Spot-winged Tit The Coal Tit, Periparus ater, is a passerine bird in the tit family Paridae. It is a...

, and probably Marsh Tit
Marsh Tit
The Marsh Tit Poecile palustris is a passerine bird in the tit family Paridae and genus Poecile, closely related to the Willow, Père David's and Songar Tits. It is small with a black crown and nape, pale cheeks, brown back and greyish-brown wings and tail. Between 8 and 11 subspecies are recognised...

.

Subspecies

There are currently 14 recognised subspecies of Great Tit:
  • P. m. newtoni, described by Pražák in 1894, is found across the British Isles
    British Isles
    The British Isles are a group of islands off the northwest coast of continental Europe that include the islands of Great Britain and Ireland and over six thousand smaller isles. There are two sovereign states located on the islands: the United Kingdom of Great Britain and Northern Ireland and...

    .
  • P. m. major, described by Linnaeus in 1758, is found throughout much of Europe, Asia Minor
    Anatolia
    Anatolia is a geographic and historical term denoting the westernmost protrusion of Asia, comprising the majority of the Republic of Turkey...

    , northern and eastern Kazakhstan
    Kazakhstan
    Kazakhstan , officially the Republic of Kazakhstan, is a transcontinental country in Central Asia and Eastern Europe. Ranked as the ninth largest country in the world, it is also the world's largest landlocked country; its territory of is greater than Western Europe...

    , southern Siberia
    Siberia
    Siberia is an extensive region constituting almost all of Northern Asia. Comprising the central and eastern portion of the Russian Federation, it was part of the Soviet Union from its beginning, as its predecessor states, the Tsardom of Russia and the Russian Empire, conquered it during the 16th...

     and northern Mongolia
    Mongolia
    Mongolia is a landlocked country in East and Central Asia. It is bordered by Russia to the north and China to the south, east and west. Although Mongolia does not share a border with Kazakhstan, its western-most point is only from Kazakhstan's eastern tip. Ulan Bator, the capital and largest...

    , as far as the mid-Amur Valley.
  • P. m. excelsus, described by Buvry in 1857, is found in northwestern Africa
    Africa
    Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...

    .
  • P. m. corsus, described by Kleinschmidt
    Otto Kleinschmidt
    Otto Kleinschmidt was a German theologist, pastor and ornithologist. He introduced a typological species concept into German ornithology. His Formenkreis theory influenced the early ideas of Erwin Stresemann. Others have considered him one of the first biogeographers...

     in 1903, is found in Portugal
    Portugal
    Portugal , officially the Portuguese Republic is a country situated in southwestern Europe on the Iberian Peninsula. Portugal is the westernmost country of Europe, and is bordered by the Atlantic Ocean to the West and South and by Spain to the North and East. The Atlantic archipelagos of the...

    , southern Spain
    Spain
    Spain , officially the Kingdom of Spain languages]] under the European Charter for Regional or Minority Languages. In each of these, Spain's official name is as follows:;;;;;;), is a country and member state of the European Union located in southwestern Europe on the Iberian Peninsula...

    , and Corsica
    Corsica
    Corsica is an island in the Mediterranean Sea. It is located west of Italy, southeast of the French mainland, and north of the island of Sardinia....

    .
  • P. m. mallorcae, described by von Jordans in 1913, is found in the Balearic Islands
    Balearic Islands
    The Balearic Islands are an archipelago of Spain in the western Mediterranean Sea, near the eastern coast of the Iberian Peninsula.The four largest islands are: Majorca, Minorca, Ibiza and Formentera. The archipelago forms an autonomous community and a province of Spain with Palma as the capital...

    .
  • P. m. ecki, described by von Jordans in 1970, is found on Sardinia
    Sardinia
    Sardinia is the second-largest island in the Mediterranean Sea . It is an autonomous region of Italy, and the nearest land masses are the French island of Corsica, the Italian Peninsula, Sicily, Tunisia and the Spanish Balearic Islands.The name Sardinia is from the pre-Roman noun *sard[],...

    .
  • P. m. niethammeri, described by von Jordans in 1970, is found on Crete
    Crete
    Crete is the largest and most populous of the Greek islands, the fifth largest island in the Mediterranean Sea, and one of the thirteen administrative regions of Greece. It forms a significant part of the economy and cultural heritage of Greece while retaining its own local cultural traits...

    .
  • P. m. aphrodite, described by Madarász in 1901, is found in southern Italy
    Italy
    Italy , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...

    , southern Greece
    Greece
    Greece , officially the Hellenic Republic , and historically Hellas or the Republic of Greece in English, is a country in southeastern Europe....

    , Cyprus
    Cyprus
    Cyprus , officially the Republic of Cyprus , is a Eurasian island country, member of the European Union, in the Eastern Mediterranean, east of Greece, south of Turkey, west of Syria and north of Egypt. It is the third largest island in the Mediterranean Sea.The earliest known human activity on the...

     and the Aegean Islands
    Aegean Islands
    The Aegean Islands are the group of islands in the Aegean Sea, with mainland Greece to the west and north and Turkey to the east; the island of Crete delimits the sea to the south, those of Rhodes, Karpathos and Kasos to the southeast...

    .
  • P. m. terrasanctae was described by Hartert
    Ernst Hartert
    Ernst Johann Otto Hartert was a German ornithologist. Hartert was born in Hamburg. He was employed by Lionel Walter Rothschild as ornithological curator of his private museum at Tring from 1892 to 1929....

     in 1910. It is found in Lebanon
    Lebanon
    Lebanon , officially the Republic of LebanonRepublic of Lebanon is the most common term used by Lebanese government agencies. The term Lebanese Republic, a literal translation of the official Arabic and French names that is not used in today's world. Arabic is the most common language spoken among...

    , Israel
    Israel
    The State of Israel is a parliamentary republic located in the Middle East, along the eastern shore of the Mediterranean Sea...

    , Jordan
    Jordan
    Jordan , officially the Hashemite Kingdom of Jordan , Al-Mamlaka al-Urduniyya al-Hashemiyya) is a kingdom on the East Bank of the River Jordan. The country borders Saudi Arabia to the east and south-east, Iraq to the north-east, Syria to the north and the West Bank and Israel to the west, sharing...

     and Syria
    Syria
    Syria , officially the Syrian Arab Republic , is a country in Western Asia, bordering Lebanon and the Mediterranean Sea to the West, Turkey to the north, Iraq to the east, Jordan to the south, and Israel to the southwest....

    .
  • P. m. karelini, described by Zarundy in 1910, is found in southeastern Azerbaijan
    Azerbaijan
    Azerbaijan , officially the Republic of Azerbaijan is the largest country in the Caucasus region of Eurasia. Located at the crossroads of Western Asia and Eastern Europe, it is bounded by the Caspian Sea to the east, Russia to the north, Georgia to the northwest, Armenia to the west, and Iran to...

     and northwestern Iran
    Iran
    Iran , officially the Islamic Republic of Iran , is a country in Southern and Western Asia. The name "Iran" has been in use natively since the Sassanian era and came into use internationally in 1935, before which the country was known to the Western world as Persia...

    .
  • P. m. blandfordi was described by Pražák in 1894. It is found in north central and southwestern Iran
    Iran
    Iran , officially the Islamic Republic of Iran , is a country in Southern and Western Asia. The name "Iran" has been in use natively since the Sassanian era and came into use internationally in 1935, before which the country was known to the Western world as Persia...

    .
  • P. m. bokharensis was described by Lichtenstein
    Martin Lichtenstein
    Martin Hinrich Carl Lichtenstein was a German physician, explorer, zoologist, and herpetologist.-Biography:...

     in 1823. It is found in southern Kazakhstan
    Kazakhstan
    Kazakhstan , officially the Republic of Kazakhstan, is a transcontinental country in Central Asia and Eastern Europe. Ranked as the ninth largest country in the world, it is also the world's largest landlocked country; its territory of is greater than Western Europe...

    , Uzbekistan
    Uzbekistan
    Uzbekistan , officially the Republic of Uzbekistan is a doubly landlocked country in Central Asia and one of the six independent Turkic states. It shares borders with Kazakhstan to the west and to the north, Kyrgyzstan and Tajikistan to the east, and Afghanistan and Turkmenistan to the south....

    , Turkmenistan
    Turkmenistan
    Turkmenistan , formerly also known as Turkmenia is one of the Turkic states in Central Asia. Until 1991, it was a constituent republic of the Soviet Union, the Turkmen Soviet Socialist Republic . Turkmenistan is one of the six independent Turkic states...

     and far north of Iran
    Iran
    Iran , officially the Islamic Republic of Iran , is a country in Southern and Western Asia. The name "Iran" has been in use natively since the Sassanian era and came into use internationally in 1935, before which the country was known to the Western world as Persia...

     and Afghanistan
    Afghanistan
    Afghanistan , officially the Islamic Republic of Afghanistan, is a landlocked country located in the centre of Asia, forming South Asia, Central Asia and the Middle East. With a population of about 29 million, it has an area of , making it the 42nd most populous and 41st largest nation in the world...

    . Was, along with following two subspecies, once treated as separate species.
  • P. m. turkestanicus, was described by Zarudny & Loudon
    John Claudius Loudon
    John Claudius Loudon was a Scottish botanist, garden and cemetery designer, author and garden magazine editor.-Background:...

     in 1905, and ranges from east Kazakhstan to extreme north west China and west Mongolia
    Mongolia
    Mongolia is a landlocked country in East and Central Asia. It is bordered by Russia to the north and China to the south, east and west. Although Mongolia does not share a border with Kazakhstan, its western-most point is only from Kazakhstan's eastern tip. Ulan Bator, the capital and largest...

    .
  • P. m. ferghanensis, was described by Buturlin
    Sergei Aleksandrovich Buturlin
    Sergei Aleksandrovich Buturlin was a Russian ornithologist.A scion of one of the oldest families of Russian nobility, Buturlin spent most his life in Russia. He went to school in Simbirsk and studied jurisprudence in St...

     in 1912, and is found in Tajikistan
    Tajikistan
    Tajikistan , officially the Republic of Tajikistan , is a mountainous landlocked country in Central Asia. Afghanistan borders it to the south, Uzbekistan to the west, Kyrgyzstan to the north, and China to the east....

     and Kyrgyzstan
    Kyrgyzstan
    Kyrgyzstan , officially the Kyrgyz Republic is one of the world's six independent Turkic states . Located in Central Asia, landlocked and mountainous, Kyrgyzstan is bordered by Kazakhstan to the north, Uzbekistan to the west, Tajikistan to the southwest and China to the east...

    .

Description

The Great Tit is large for a tit at 12.5 to 14 cm (4.9 to 5.5 in) in length, and has a distinctive appearance that makes it easy to recognise. The nominate race P. major major has a bluish-black crown
Crown (anatomy)
A crown is the top of the head.The following birds and other animals are said to have a crown on their head:* Cranes** Grey-crowned Crane** Red-crowned Crane** Black-crowned Crane* Crowned eagle* Gray-crowned Rosy Finch* Yellow-crowned Gonolek...

, black neck, throat, bib and head, and white cheeks and ear coverts. The breast is bright lemon-yellow and there is a broad black mid-line stripe running from the bib to vent. There is a dull white spot on the neck turning to greenish yellow on the upper nape. The rest of the nape and back are green tinged with olive. The wing-coverts
Covert (feather)
A covert feather on a bird is one of a set of feathers, called coverts, which as the name implies, cover other feathers. The coverts help to smooth airflow over the wings and tail.- Wing-coverts :...

 are green, the rest of the wing is bluish-grey with a white-wing-bar. The tail is bluish grey with white outer tips. The plumage
Plumage
Plumage refers both to the layer of feathers that cover a bird and the pattern, colour, and arrangement of those feathers. The pattern and colours of plumage vary between species and subspecies and can also vary between different age classes, sexes, and season. Within species there can also be a...

 of the female is similar to that of the male except that the colours are overall duller; the bib is less intensely black, as is the line running down the belly, which is also narrower and sometimes broken. Young birds are like the female, except that they have dull olive-brown napes and necks, greyish rumps, and greyer tails, with less defined white tips.
There is some variation in the subspecies. P. m. newtoni is like the nominate race but has a slightly longer bill, the mantle is slightly deeper green, there is less white on the tail tips, and the ventral mid-line stripe is broader on the belly. P. m. corsus also resembles the nominate form but has duller upperparts, less white in the tail and less yellow in the nape. P. m. mallorcae is like the nominate subspecies, but has a larger bill, greyer-blue upperparts and slightly paler underparts. P. m. ecki is like P. m. mallorcae except with bluer upperparts and paler underparts. P. m. excelsus is similar to the nominate race but has much brighter green upperparts, bright yellow underparts and no (or very little) white on the tail. P. m. aphrodite has darker, more olive-grey upperparts, and the underparts are more yellow to pale cream. P. m. niethammeri is similar to P. m. aphrodite but the upperparts are duller and less green, and the underparts are pale yellow. P. m. terrasanctae resembles the previous two subspecies but has slightly paler upperparts. P. m. blandfordi is like the nominate but with a greyer mantle and scapulars and pale yellow underparts, and P. m. karelini is intermediate between the nominate and P. m. blandfordi, and lacks white on the tail. The plumage of P. m. bokharensis is much greyer, pale creamy white to washed out grey underparts, a larger white cheep patch, a grey tail, wings, back and nape. It is also slightly smaller, with a smaller bill but longer tail. The situation is similar for the two related subspecies in the Turkestan Tit group. P. m. turkestanicus is like P. m. bokharensis but with a larger bill and darker upperparts. P. m. ferghanensis is like P. m. bokharensis but with a smaller bill, darker grey on the flanks and a more yellow wash on the juvenile birds.

The colour of the male bird's breast has been shown to correlate with stronger sperm, and is one way that the male demonstrates his reproductive superiority to females. Higher levels of carotenoid
Carotenoid
Carotenoids are tetraterpenoid organic pigments that are naturally occurring in the chloroplasts and chromoplasts of plants and some other photosynthetic organisms like algae, some bacteria, and some types of fungus. Carotenoids can be synthesized fats and other basic organic metabolic building...

 increase the intensity of the yellow of the breast its colour, and also enable the sperm to better withstand the onslaught of free radical
Radical (chemistry)
Radicals are atoms, molecules, or ions with unpaired electrons on an open shell configuration. Free radicals may have positive, negative, or zero charge...

s. Carotenoids cannot be synthesized by the bird and have to be obtained from food, so a bright colour in a male demonstrates his ability to obtain good nutrition. The width of the male's ventral stripe, which varies with individual, is selected for by females, with higher quality females apparently selecting males with wider stripes.

Voice

The Great Tit is, like other tits, a vocal bird, and has up to 40 types of calls and songs. The calls are generally the same between the sexes, but the male is much more vocal and the female rarely calls. Soft single notes such as "pit", "spick", or "chit" are used as contact calls. A loud "tink" is used by adult males as an alarm or in territorial disputes. One of the most familiar is a "teacher, teacher", often likened to a squeaky wheelbarrow
Wheelbarrow
A wheelbarrow is a small hand-propelled vehicle, usually with just one wheel, designed to be pushed and guided by a single person using two handles to the rear, or by a sail to push the ancient wheelbarrow by wind. The term "wheelbarrow" is made of two words: "wheel" and "barrow." "Barrow" is a...

 wheel, which is used in proclaiming ownership of a territory. In former times, English folk considered the "saw-sharpening" call to be a foretelling of rain. There is little geographic variation in calls, but tits from the two south Asian groups recently split from the Great Tit do not recognise or react to the calls of the temperate Great Tits.

One explanation for the Great Tit’s wide repertoire is the Beau Geste
Beau Geste
Beau Geste is a 1924 adventure novel by P. C. Wren. It has been adapted for the screen several times.-Plot summary:Michael "Beau" Geste is the protagonist. The main narrator , by contrast, is his younger brother John...

 theory. The eponymous hero of the novel propped dead soldiers against the battlements to give the impression that his fort was better defended than was really the case. Similarly, the multiplicity of calls gives the impression that the tit’s territory is more densely occupied than it actually is. Whether the theory is correct or not, those birds with large vocabularies are socially dominant and breed more successfully.

Distribution, movements and habitat

The Great Tit has a wide distribution across much of Eurasia. It is found across all of Europe except for Iceland and northern Scandinavia, including numerous Mediterranean islands. In North Africa it is found in Morocco, Algeria and Tunisia. It also occurs across the Middle East
Middle East
The Middle East is a region that encompasses Western Asia and Northern Africa. It is often used as a synonym for Near East, in opposition to Far East...

, and parts of central Asia from northern Iran and Afghanistan to Mongolia
Mongolia
Mongolia is a landlocked country in East and Central Asia. It is bordered by Russia to the north and China to the south, east and west. Although Mongolia does not share a border with Kazakhstan, its western-most point is only from Kazakhstan's eastern tip. Ulan Bator, the capital and largest...

, as well as across northern Asia from the Urals
Ural Mountains
The Ural Mountains , or simply the Urals, are a mountain range that runs approximately from north to south through western Russia, from the coast of the Arctic Ocean to the Ural River and northwestern Kazakhstan. Their eastern side is usually considered the natural boundary between Europe and Asia...

 as far east as northern China and the Amur Valley.

The Great Tit occupies a range of habitats. It is most commonly found in open deciduous woodland, mixed forests and forest edges. In dense forests, including conifer
Pinophyta
The conifers, division Pinophyta, also known as division Coniferophyta or Coniferae, are one of 13 or 14 division level taxa within the Kingdom Plantae. Pinophytes are gymnosperms. They are cone-bearing seed plants with vascular tissue; all extant conifers are woody plants, the great majority being...

 forests it is usually found in forest clearings. In northern Siberia
Siberia
Siberia is an extensive region constituting almost all of Northern Asia. Comprising the central and eastern portion of the Russian Federation, it was part of the Soviet Union from its beginning, as its predecessor states, the Tsardom of Russia and the Russian Empire, conquered it during the 16th...

 it is found in boreal taiga
Taiga
Taiga , also known as the boreal forest, is a biome characterized by coniferous forests.Taiga is the world's largest terrestrial biome. In North America it covers most of inland Canada and Alaska as well as parts of the extreme northern continental United States and is known as the Northwoods...

. In North Africa it prefers oak
Oak
An oak is a tree or shrub in the genus Quercus , of which about 600 species exist. "Oak" may also appear in the names of species in related genera, notably Lithocarpus...

 forests as well as stands of Atlas cedar
Atlas Cedar
Cedrus atlantica, the Atlas Cedar, is a cedar native to the Atlas Mountains of Algeria and Morocco . A majority of the modern sources treat it as a distinct species Cedrus atlantica, but some sources consider it a subspecies of Lebanon Cedar Cedrus atlantica, the Atlas Cedar, is a cedar native to...

 and even palm groves. In the east of its range in Siberia, Mongolia and China it favours riverine willow and birch forest. Riverine woodlands of willows, poplars are among the habitats of the Turkestan group in central Asia, as well as low scrubland, oases; at higher altitudes it occupies habitats ranging from dense deciduous and coniferous forests to open areas with scattered trees.

The Great Tit is generally not migratory
Bird migration
Bird migration is the regular seasonal journey undertaken by many species of birds. Bird movements include those made in response to changes in food availability, habitat or weather. Sometimes, journeys are not termed "true migration" because they are irregular or in only one direction...

. Pairs will usually remain near or in their territory year round, even in northern parts of their range. Young birds will disperse from their parents' territory, but usually not far. Populations may become irruptive in poor or harsh winters, meaning that groups of up to a thousand birds will may unpredictably move from northern Europe to the Baltic, the Netherlands, Britain and even as far as the southern Balkans.

The Great Tit was unsuccessfully introduced into the United States; birds were set free near Cincinnati, Ohio between 1872 and 1874 but failed to become established. Suggestions that they were an excellent control measure for codling moth
Codling moth
The codling moth is a member of the Lepidopteran family Tortricidae. They are known as an agricultural pest, their larva being the common apple worm or maggot. It is native to Europe and was introduced to North America, where it has become one of the regular pests of apple orchards. It is found...

s nearly led to their introduction to some new areas particularly in the United States of America, however this plan was not implemented. Birds were later introduced to the Almaty Province
Almaty Province
Almaty is a province of Kazakhstan. Its capital is Taldykorgan, which has a population of 118,400. The province itself has 1,603,700 people.-Geography:...

 in what is now Kazakhstan
Kazakhstan
Kazakhstan , officially the Republic of Kazakhstan, is a transcontinental country in Central Asia and Eastern Europe. Ranked as the ninth largest country in the world, it is also the world's largest landlocked country; its territory of is greater than Western Europe...

 in 1960–61 and became established, although their present status is unclear.

Diet and feeding

Great Tits are primarily insectivorous
Insectivore
An insectivore is a type of carnivore with a diet that consists chiefly of insects and similar small creatures. An alternate term is entomophage, which also refers to the human practice of eating insects....

 in the summer, feeding on insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s and spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...

s which they capture by foliage
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

 gleaning
Gleaning (birds)
Gleaning is a term for a feeding strategy by birds in which they catch invertebrate prey, mainly arthropods, by plucking them from foliage or the ground, from crevices such as rock faces and under the eaves of houses, or even, as in the case of ticks and lice, from living animals. This behavior is...

. Invertebrate prey that are taken include cockroaches, grasshoppers and crickets, lacewings, earwigs, bugs (Hemiptera), ants, flies (Diptera), caddis flies, beetles, scorpion flies, woodlice, harvestmen, bees and wasps, snails and woodlice. During the breeding season, the tits prefer to feed protein-rich caterpillars to their young. A study published in 2007 found that Great Tits helped to reduce caterpillar damage in apple orchards by as much as 50%. Nestlings also undergo a period in their early development where they are fed a number of spiders, possibly for nutritional reasons. In autumn and winter, when insect prey becomes scarcer, Great Tits add berries
Berry
The botanical definition of a berry is a fleshy fruit produced from a single ovary. Grapes are an example. The berry is the most common type of fleshy fruit in which the entire ovary wall ripens into an edible pericarp. They may have one or more carpels with a thin covering and fleshy interiors....

 and seed
Seed
A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...

s to their diet. Seeds and fruit usually come from deciduous trees and shrubs, and include the seeds of beech and hazel. Where it is available they will readily take table scraps, peanuts and sunflower seeds from bird tables. In particularly severe winters they may consume 44% of their body weight in sunflower seeds. They often forage on the ground, particularly in years with high beech
Beech
Beech is a genus of ten species of deciduous trees in the family Fagaceae, native to temperate Europe, Asia and North America.-Habit:...

 mast
Mast (botany)
Mast is the "fruit of forest trees like acorns and other nuts". It is also defined as "the fruit of trees such as beech, and other forms of Cupuliferae". Alternatively, it can also refer to "a heap of nuts"....

 production. Great Tits, along with other tits, will join winter mixed-species foraging flocks.

Large food items, such as large seeds or prey, are dealt with by "hold-hammering", where the item is held with one or both feet and then struck with the bill until it is ready to eat. Using this method, a Great Tit can get into a hazelnut
Hazelnut
A hazelnut is the nut of the hazel and is also known as a cob nut or filbert nut according to species. A cob is roughly spherical to oval, about 15–25 mm long and 10–15 mm in diameter, with an outer fibrous husk surrounding a smooth shell. A filbert is more elongated, being about twice...

 in about twenty minutes. When feeding young, adults will hammer off the heads off large insects to make them easier to consume, and remove the gut from caterpillars so that the tannin
Tannin
A tannin is an astringent, bitter plant polyphenolic compound that binds to and precipitates proteins and various other organic compounds including amino acids and alkaloids.The term tannin refers to the use of...

s in the gut will not retard the chick's growth.

Great Tits combine dietary versatility with a considerable amount of intelligence and the ability to solve problems with insight learning, that is to solve a problem through insight rather than trial and error. In England, Great Tits learned to break the foil caps of milk bottles delivered at the doorstep of homes to obtain the cream at the top. This behaviour, first noted in 1921, spread rapidly in the next two decades. In 2009, Great Tits were reported killing and eating pipistrelle
Pipistrellus
Pipistrellus is a genus of bats in the family Vespertilionidae and subfamily Vespertilioninae. The name of the genus is derived from the Italian word Pipistrello, meaning "bat"....

 bats. This is the first time a songbird has been seen to hunt bats. The tits only do this during winter when the bats are hibernating and other food is scarce. They have also been recorded using tools
Tool use by animals
Tools are used by some animals, particularly primates, to perform simple tasks such as the acquisition of food, or grooming. Originally thought to be a skill only possessed by humans, tool use requires some level of intelligence. Primates have been observed exploiting sticks and stones to...

, using a conifer needle in the bill to extract larvae from a hole in a tree.

Breeding

Great Tits are monogamous
Monogamy
Monogamy /Gr. μονός+γάμος - one+marriage/ a form of marriage in which an individual has only one spouse at any one time. In current usage monogamy often refers to having one sexual partner irrespective of marriage or reproduction...

 breeders and establish breeding territories
Territory (animal)
In ethology the term territory refers to any sociographical area that an animal of a particular species consistently defends against conspecifics...

. These territories are established in late January and defence begins in late winter or early spring. Territories are usually reoccupied in successive years, even if one of the pair dies, so long as the brood is raised successfully. Females are likely to disperse to new territories if their nest is predated the previous year. If the pair divorces for some reason then the birds will disperse, with females travelling further than males to establish new territories. Although the Great Tit is socially monogamous, extra-pair
Promiscuity
In humans, promiscuity refers to less discriminating casual sex with many sexual partners. The term carries a moral or religious judgement and is viewed in the context of the mainstream social ideal for sexual activity to take place within exclusive committed relationships...

 copulations are frequent. One study in Germany found that 40% of nests contained some offspring fathered by parents other than the breeding male and that 8.5% of all chicks were the result of cuckold
Cuckold
Cuckold is a historically derogatory term for a man who has an unfaithful wife. The word, which has been in recorded use since the 13th century, derives from the cuckoo bird, some varieties of which lay their eggs in other birds' nests...

ry.

Great Tits are seasonal breeders. The exact timing of breeding varies by a number of factors, most importantly location. Most breeding occurs between January and September; in Europe the breeding season usually begins after March. In Israel there are exceptional records of breeding during the months of October to December. The amount of sunlight and daytime temperatures will also affect breeding timing. One study found a strong correlation between the timing of laying and the peak abundance of caterpillar prey, which is in turn correlated to temperature. On an individual level, younger females tend to start laying later than older females.
Great Tits are cavity nesters, breeding in a hole that is usually inside a tree, although occasionally in a wall or rock face, and they will readily take to nest box
Nest box
A nest box, also spelled nestbox is a man-made box provided for animals to nest in. Nest boxes are most frequently utilized for wild and domesticated birds, in which case they are also called birdhouses, but some mammalian species may also use them. Birdhouses are the most common types of nest...

es. The nest inside the cavity is built by the female, and is made of plant fibres, grasses, moss, hair, wool and feathers. The number in the clutch
Clutch (eggs)
A clutch of eggs refers to all the eggs produced by birds or reptiles, often at a single time, particularly those laid in a nest.In birds, destruction of a clutch by predators, , results in double-clutching...

 is often very large, as many as 18, but five to twelve is more common. Clutch size is smaller when birds start laying later, and is also lower when the density of competitors is higher. Second broods tend to have smaller clutches. Insularity also affects clutch size, with Great Tits on offshore islands laying smaller clutches with larger eggs than mainland birds. The eggs are white with red spots. The female undertakes all incubation
Avian incubation
Incubation refers to the process by which certain oviparous animals hatch their eggs, and to the development of the embryo within the egg. The most vital factor of incubation is the constant temperature required for its development over a specific period. Especially in domestic fowl, the act of...

 duties, and is fed by the male during incubation. The bird is a close sitter, hissing when disturbed. The timing of hatching, which is best synchronised with peak availability of prey, can be manipulated when environmental conditions change after the laying of the first egg by delaying the beginning of incubation, laying more eggs or pausing during incubation. The incubation period is between 12 to 15 days.

The chicks, like those of all tits, are hatched unfeathered and blind. Once feathers begin to erupt, the nestlings are unusual for altricial
Altricial
Altricial, meaning "requiring nourishment", refers to a pattern of growth and development in organisms which are incapable of moving around on their own soon after hatching or being born...

 birds in having plumage coloured with carotenoid
Carotenoid
Carotenoids are tetraterpenoid organic pigments that are naturally occurring in the chloroplasts and chromoplasts of plants and some other photosynthetic organisms like algae, some bacteria, and some types of fungus. Carotenoids can be synthesized fats and other basic organic metabolic building...

s similarly to their parents (in most species it is dun-coloured to avoid predation). The nape
Nape
The nape is the back of the neck. In technical anatomical/medical terminology, the nape is referred to by the word nucha, which also gives the adjective corresponding to "nape" in English, "nuchal"....

 is yellow and attracts the attention of the parents by its ultraviolet
Ultraviolet
Ultraviolet light is electromagnetic radiation with a wavelength shorter than that of visible light, but longer than X-rays, in the range 10 nm to 400 nm, and energies from 3 eV to 124 eV...

 reflectance. This may be to make them easier to find in low light, or be a signal of fitness
Fitness (biology)
Fitness is a central idea in evolutionary theory. It can be defined either with respect to a genotype or to a phenotype in a given environment...

 to win the parents' attention. This patch turns white after the first moult at age two months, and diminishes in size as the bird grows.

Chicks are fed by both parents, usually receiving 6 to 7 g (0.211643772630672 to 0.246917734735784 oz) of food a day. Both parents provision the chicks with food and aid in nest sanitation by removing faecal packets, with no difference in the feeding effort between the sexes. The nestling period is between 16 to 22 days, with chicks being independent of the parents eight days after fledging
Fledge
Fledge is the stage in a young bird's life when the feathers and wing muscles are sufficiently developed for flight. It also describes the act of a chick's parents raising it to a fully grown state...

. Feeding of the fledgeling may continue after independence, lasting up to 25 days in chicks from the first brood, but as long as 50 days in the second brood. Nestlings from second broods have weaker immune systems and body condition than those from first broods, and hence have a lower juvenile survival rate.

Ecology

The Eurasian Sparrowhawk is a predator of Great Tits, with the young from second broods being at higher risk partly because of the hawk’s greater need for food for its own developing young. The nests of Great Tits are raided by Great Spotted Woodpecker
Great Spotted Woodpecker
The Great Spotted Woodpecker , Dendrocopos major, is a bird species of the woodpecker family . It is distributed throughout Europe and northern Asia, and usually resident year-round except in the colder parts of its range...

s, particularly when nesting in certain types of nest boxes. Other nest predators include introduced
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...

 Grey Squirrels
Eastern Gray Squirrel
The eastern gray squirrel is a tree squirrel in the genus Sciurus native to the eastern and midwestern United States, and to the southerly portions of the eastern provinces of Canada...

 (in Britain) and Least Weasel
Least Weasel
The least weasel is the smallest member of the Mustelidae , native to Eurasia, North America and North Africa, though it has been introduced elsewhere. It is classed as Least Concern by the IUCN, due to its wide distribution and presumably large population...

s, which are able to take nesting adults as well. A species of biting louse (Mallophaga
Chewing louse
Mallophaga is a suborder of lice, known as chewing lice, biting lice or bird lice, containing more than 3000 species. They have paurometabolis or incomplete metamorphosis....

) described as Rostrinirmus hudeci was isolated and described in 1981 from Great Tits in central Europe. The hen flea Ceratophyllus gallinae is exceedingly common in the nests of Blue and Great Tits. It was originally a specialist tit flea, but the dry, crowded conditions of chicken runs enabled it to flourish with its new host. This flea is preferentially predated by the clown beetle Gnathoncus punctulatus, The rove beetle
Rove beetle
The rove beetles are a large family of beetles, primarily distinguished by their short elytra that leave more than half of their abdomens exposed. With over 46,000 species in thousands of genera, the group is the second largest family of beetles after the Curculionidae...

 Microglotta pulla also feeds on fleas and their larvae. Although these beetles often remain in deserted nests, they can only breed in the elevated temperatures produced by brooding birds, tits being the preferred hosts.

Relationship with humans

The Great Tit is a popular garden bird due to its acrobatic performances when feeding on nuts or seed. Its willingness to move into nest boxes has made it a valuable study subject in ornithology
Ornithology
Ornithology is a branch of zoology that concerns the study of birds. Several aspects of ornithology differ from related disciplines, due partly to the high visibility and the aesthetic appeal of birds...

, and it is one of the best studied birds in the world. It has been particularly useful as a model for the study of the evolution of various life-history traits, particularly clutch size. A study of a literature database search
Biological Abstracts
Biological Abstracts is a database produced by Thomson Reuters through its subsidiary BIOSIS. It includes abstracts from peer-reviewed academic journal articles in the fields of biology, biochemistry, biotechnology, botany, pre-clinical and experimental medicine, pharmacology, zoology, agriculture,...

 found 1,349 articles relating to Parus major for the period between 1969 and 2002.

The Great Tit has generally adjusted to human modifications of the environment. It remains commoner, and has better breeding success, in areas with undisturbed forest cover, but it has adapted to human modified habitats, including urban areas, where it can be very common. For example, the breeding population in the city of Sheffield
Sheffield
Sheffield is a city and metropolitan borough of South Yorkshire, England. Its name derives from the River Sheaf, which runs through the city. Historically a part of the West Riding of Yorkshire, and with some of its southern suburbs annexed from Derbyshire, the city has grown from its largely...

 (a city of half a million people) has been estimated at 17,164 individuals. In adapting to human environments its song has been observed to change in noise-polluted urban environments. In areas with low frequency background noise pollution, the song has a higher frequency than in quieter areas. This tit has expanded its range, moving northwards into Scandinavia and Scotland, and south into Israel and Egypt. The total population is estimated at between 300–1,100 million birds in a range of 32.4 million km2 (12.5 million mi2). While there have been some localised declines in population in areas with poorer quality habitats, its large range and high numbers mean that the Great Tit is not considered to be threatened, and it is classed as Least Concern
Least Concern
Least Concern is an IUCN category assigned to extant taxon or lower taxa which have been evaluated but do not qualify for any other category. As such they do not qualify as threatened, Near Threatened, or Conservation Dependent...

 on the IUCN Red List
IUCN Red List
The IUCN Red List of Threatened Species , founded in 1963, is the world's most comprehensive inventory of the global conservation status of biological species. The International Union for Conservation of Nature is the world's main authority on the conservation status of species...

.

External links

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